The oldest psyllipsocid booklice, in Lower Cretaceous amber from Lebanon (Psocodea, Trogiomorpha, Psocathropetae, Psyllipsocidae)

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ZooKeys 130: 153-165 (2011)

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The oldest psyllipsocid booklice, in Lower Cretaceous
amber from Lebanon (Psocodea, Irogiomorpha,
Psocathropetae, Psyllipsocidae)

Dany Azar'', André Nel*#

| Lebanese University, Faculty of Sciences II, Department of Natural Sciences, Fanar, Fanar — Matn — PO.
Box 26110217, Lebanon 2 CNRS UMR 7205, CP 50, Entomologie, Muséum National d'Histoire Naturelle,
45 rue Buffon, F-75005, Paris, France

T urn:lsid:zoobank. org:author:2F73 1869-E5AA-49A5-8806-3ACF2211134B
¢ urn:lsid:zoobank.org:author:98 DF555A-16A0-4073-871 C-E38BB506C676

Corresponding author: Dany Azar (azar@mnhn.fr)

Academic editor: D. Shcherbakov | Received 24 April 2011 | Accepted 4 July 2011 | Published 24 September 2011
urn:lsid:zoobank.org:pub:A5D2FFAD-BB72-4CF3-A089-CF4FC5BED785

Citation: Azar D, Nel A (2011) The oldest psyllipsocid booklice, in Lower Cretaceous amber from Lebanon (Psocodea,
Trogiomorpha, Psocathropetae, Psyllipsocidae). In: Shcherbakov DE, Engel MS, Sharkey MJ (Eds) Advances in the
Systematics of Fossil and Modern Insects: Honouring Alexandr Rasnitsyn. ZooKeys 130: 153-165. doi: 10.3897/
zookeys. 130.1430

Abstract
Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., of Psyllipsocidae is described and figured from the
Lower Cretaceous amber of Lebanon. The position of the new taxon is discussed and the fossil is com-

pared to other psyllipsocids. The species represents the earliest record of the family Psyllipsocidae.

Keywords

New genus, new species, Psyllipsocidae, Lower Cretaceous, Lebanese amber

Introduction

The Psocodea (Psocoptera + Phthiraptera) is a rather small order with about 10,000
valid extant species. Earlier, fossils from the Permian were assigned to the Psocodea but
the attribution of these is uncertain. The first unquestionable records of psocodeans
are from the Middle Jurassic of China (Huang et al. 2008). The Psyllipsocidae are a

relatively small family of trogiomorphan psocathropetan Psocodea, including about 50

Copyright Dany Azar,André Nel. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
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154 Dany Azar & André Nel / ZooKeys 130: 153-165 (2011)

extant species in five genera distributed worldwide. Psyllipsocids live in caves, on rock
surfaces, and among dead leaves. To date four fossils have been assigned to Psyllipso-
cidae: Psyllipsocus eocenicus Nel et al., 2005 (lowermost Eocene French Oise amber),
?Psyllipsocus banksi Cockerell, 1916 (mid-Cretaceous Burmese amber), a ”Psyllipsocus
sp. (nymph)“ from Lower Miocene Mexican amber (Mockford 1969), and Khatangia
inclusa Vishniakova, 1975 (Late Cretaceous Taimyr amber) but the last three attribu-
tions to Psyllipsocidae are dubious and require revision.

The Lower Cretaceous Lebanese amber is so far the oldest (Neocomian) with com-
plete macroscopic biological inclusions (Azar 2007). Its formation corresponds to the
epoch of the origin and rise of flowering plants, a critical period for the study of the
evolution, with extinction of many ancient insect groups and emergence of effectively
modern lineages. Up to now nine species of psocids have been described from Leba-
nese amber (Perrichot et al. 2003; Azar and Nel 2004; Grimaldi and Engel 2006; Azar
et al. 2008, 2010).

Herein a new psyllipsocid genus and species is described from the Lower Creta-
ceous amber of Lebanon, representing the earliest record of this family.

Material and methods

The amber piece containing the inclusion was cut, shaped, and polished, before prepa-
rations between two coverslips and in a Canada balsam medium, as described in Azar
et al. (2003). The specimen was examined with a Nikon SZ10 stereomicroscope and a
Leitz Laborlux-12 compound microscope, both equipped with camera lucida for line
drawings. Photographs were made with an Olympus FE-5000 digital camera.

The works of Smithers (1972, 1990), Lienhard (1998) and Mockford (1993) are
followed herein for the systematics of Psocodea. ‘The wing venation nomenclature and
terminology of body structures of Smithers (1972) and Lienhard (1998) were adopted.

Systematic Paleontology

Suborder Trogiomorpha Roesler, 1944
Infraorder Psocathropetae Pearman, 1936
Family Psyllipsocidae Enderlein, 1911

Libanopsyllipsocus gen. n.
urn:lsid:zoobank.org:act:E646B958-D9 1 F-4C29-9 17D-502035368507
http://species-id.net/wiki/Libanopsyllipsocus

Type species. Libanopsyllipsocus alexanderasnitsyni sp. n.
Etymology. After “Libano” (Lebanese in Latin), and “Psyllipsocus”, type genus of

the family Psyllipsocidae; gender masculine.

Oldest Psyllipsocidae from the Lebanese amber... 155

Diagnosis. Hypopharynx with two distinct filaments. Macropterous; forewing
without pterostigma; M two-branched; areola postica elongate and free; Cu2 and A
ending together forming a nodulus. Hindwing with R1 absent; Rs two-branched;
M simple; basal cell present. Distal end of tibia with two inner spines; tarsi three-
segmented; distal pretarsi with claw with preapical tooth, and without pulvillus and
microtrichia.

Libanopsyllipsocus alexanderasnitsyni, sp. n.
urn:lsid:zoobank.org:act:F79BD9BC-BB80-44B6-AEA4-99748A240347
http://species-id.net/wiki/Libanopsyllipsocus_alexanderasnitsyni

Figs 1-12

Holotype. Specimen n° 30 (male, coll. Azar); provisionally deposited in the Muséum
National d’ Histoire Naturelle, Paris, France.

Locality and horizon. Lower Cretaceous, Upper Barremian — Lower Aptian, am-
ber of Hammana-Mdeyrij, Caza, Baabda, Mohafazat Jabal Loubnan, central Lebanon.

Etymology. In honor of our friend Prof. Alexandr P. Rasnitsyn, a world authority
in entomology and palaeoentomology.

Diagnosis. As for the genus (vide supra).

Description. Dorsal parts of head and body not preserved. Body and wings with-
out scales (Figs 1-2). Head elongate (but cannot be measured accurately due to state
of preservation), sublingual sclerite and hypopharynx with two distinct filaments (Fig.
3). Forewing transparent (Fig. 4), with infrequent small setae, 0.96 mm long, 0.35
mm broad; apex rounded; Sc badly preserved, not reaching C nor R; R1 reaching cos-
tal margin 0.73 mm from wing base; pterostigma absent; Rs separating from R 0.46
mm from wing base; Rs bifurcating into R2+3 and R4+5 at 0.76 mm; R2+3 curved,
reaching wing margin 0.83 mm distally; R4+5 curved, reaching wing margin at 0.95
mm distally; a transverse crossvein between Rs and M; fork of M into M1 and M2 0.8
mm from wing base, but one wing is aberrant in having M simple [a frequent teratol-
ogy in Psyllipsocidae as noted by Lienhard (1998)]; Cul bifurcating into Cula and
Culb 0.36 mm from wing base forming a free elongate and narrow areola postica;
Cula much longer than Culb; Cu2 (badly preserved) joining anal vein at posterior
wing margin in a nodulus 0.28 mm from wing base (Fig. 5); A strongly convex. Wing
margin glabrous (Fig. 6), similar in structure to that illustrated by Lienhard (1998:
111, fig. 31b). Hind wing transparent (Fig. 4), bare and smaller than forewing, about
0.75 mm long, 0.24 mm broad; free or individualized and distinct arm of R1 missing;
Rs forked into R2+3 and R4+5 0.16 mm from wing apex; basal cell present; simple M
reaching wing margin apically; basal part of wing badly preserved. Hind leg with tibia
+ tarsi slightly longer than abdomen. Coxal rasp of Pearman’s organ present on hind
leg (Fig. 7), but without tympanum (the coxal rasp is preserved on one leg as the coxal
area is missing on the other leg). Tibiae with two apical spurs on inner sides; all tarsi
three-segmented with first tarsomere longer than others; pretarsal claws with a preapi-

156 Dany Azar & André Nel / ZooKeys 130: 153-165 (2011)

cal tooth, without microtrichia nor pulvillus (Fig. 8). Abdomen 0.52 mm long, 0.32
mm broad; telson with epiproct and paraproct as represented in Figs 9-12, paraproct
with a rounded hemispherical sensory area of trichobotria, and a spine in its ventral
inner side; male genitalia with simple hypandrium, aedeagus with phallosome present-
ing parameres with curved arms not fused anteriorly as represented in Fig. 9 (it can be
seen owing to transparency of sclerites and is comparable in structure to what occurs

in other Psyllipsocidae (cf. Lienhard 1998: 112, fig. 32b).

Discussion

Following the keys of Smithers (1972), Mockford (1993), and Lienhard (1998), Liban-
opsyllipsocus alexanderasnitsyni gen. et sp. n. would fall into Trogiomorpha and can be
placed in Psyllipsocidae owing to the following combination of characters, considered
diagnostic of the family (Smithers 1972, 1990; Mockford 1993; Lienhard 1998), and
present in the current fossil: head elongate; hypopharynx with two distinct sclerotized
filaments; presence of nodulus; posterior legs with tibia + tarsi longer than abdomen;
tarsi three-segmented; distal pretarsi with claw bearing preapical tooth, without pulvil-
lus; paraproct with anal spine; phallosome with two curved arms not fused anteriorly.
Other diagnostic characters of the family Psyllipsocidae could be present but cannot be
seen in the specimen owing to the state of preservation, i.e., some parts of the animal
are missing.

Our fossil presents a striking and astonishing similarity in some of the wing features
(like elongate areola postica and M 2-branched in forewing) with the psocids belong-
ing to the troctomorphan family Pachytroctidae, to the point that any psocidologist at
first glance to the specimen would assign it to this family, and as in Pachytroctidae only
females could be winged, he would tell directly that this is a female! A minute examina-
tion of the specimen shows that the fossil is a male with genitalia structure comparable
to those of Psyllipsocidae as figured by Lienhard (1998: 112, fig. 32b) with the very
particular conical phallosome directed forwards. Moreover, our fossil could not belong
to the Pachytroctidae for several reasons: (1) in the forewing Cu2 and A are meeting
together on the wing margin thus forming a nodulus, the absence of this feature being
characteristic for Pachytroctidae; (2) presence of Pearman’s coxal organ in hind legs,
here too the absence of this feature is characteristic of the Pachytroctidae (Yoshizawa
and Lienhard 2010), and even for the whole Nanopsocetae that include this family; (3)
presence of distinct separated sclerotized filaments of hypopharynx characteristic of all
Trogiomorpha, whereas in Troctomorpha (that enclose Pachytroctidae) these filaments
are fused.

Our fossil shows a derived character for its hind wing with M simple, though the
majority of the fully-winged extant species of Psyllipsocidae, like most of the trogio-
morphan psocids, have hind wings with M forked (a feature considered plesiomorphic
in general for the Psocodea).

Oldest Psyllipsocidae from the Lebanese amber... 157

Libanopsyllipsocus gen. n. differs from all modern genera by several features of its
wings; i.e., from the genera Dorypteryx Aaron, 1899 and Psocathropos Ribaga, 1899 by
the glabrous anterior margin of the forewing; from Pseudodorypteryx Garcia-Aldrete,
1984 and Dorypteryx Aaron, 1899 by the broad forewing. Libanopsyllipsocus gen. n.
resembles mostly the genus Psy/lipsocus Sélys-Longchamps, 1872 but differs from it in
its forewing without a pterostigma and M two-branched, and in its hind wing with
R1 absent and M simple. Libanopsyllipsocus gen. n. differs from the genus Khatangia
Vishniakova, 1975 in the absence of a pterostigma and an elongate areola postica in the
forewing. All these differences and many others lead us to the proposal of a new genus.

The monophyly of the family Psyllipsocidae is supported by one character of the
female genitalia: the spermathecal sac with complex scleritizations at the origin and
usually with an accessory vesicle. Molecular analysis also supports the monophyly of
the family (Yoshizawa et al. 2006). Libanopsyllipsocus alexanderasnitsyni gen. et sp. n.
represents the oldest record of the Psyllipsocidae and demonstrates that this family is
at least as old as the lowermost Cretaceous, if not older. Unexpectedly some features
of this very old fossil are probably derived, viz., the absence of a pterostigma in the
forewing and M simple in the hind wing. The discovery of new fossils belonging to
this family could bring more information on the polarity and potential homoplasy of
some characters in use for the classification, and increase as such our knowledge of the
evolutionary history of this lineage.

Acknowledgments

The authors are grateful to Michael S. Engel for his important comments on an earlier
version of the manuscript. This paper is a contribution to the team project “Biodiver-
sity: Origin, Structure, Evolution and Geology” granted to DA by Lebanese University.

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Oldest Psyllipsocidae from the Lebanese amber... 159

Plates

Figure |. Photograph of habitus of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male,

specimen number 30.

Figure 2. Drawing of habitus of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male, scale

bar = 0.3 mm.

160 Dany Azar & André Nel / ZooKeys 130: 153-165 (2011)

Figure 3. Photograph of hypopharynx filaments (arrows) of Libanopsyllipsocus alexanderasnitsyni gen. et

Rl

sp. n.. holotvve. male.

Sc R2+3

R4+5

.
eaeee centr ee,
tee,
ee

Pa M1

..
Mies
"Beesels,

M2
Cu2 Gulla

R233

R4+5

Cul Ml

Figure 4. Drawing of wings of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male, scale

bar = 0.3 mm.

Oldest Psyllipsocidae from the Lebanese amber... 161

Figure 6. Microphotograph of structure of forewing margin.

162 Dany Azar & André Nel / ZooKeys 130: 153-165 (2011)

ra

Lustiien— _

Figure 7. Microphotograph of hind leg coxal rasp (Pearman’s organ).

Oldest Psyllipsocidae from the Lebanese amber... 163

ome

Figure 8. Drawing of pretarsal claw of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male,
scale bar = 0.03 mm.

Figure 9. Drawing of aedeagus of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male; Ep
= epiproct, Hyp = hypandrium, Par = paraproct, par = paramers, Pha = phallosome, sp = anal spine, trich
= trichobothria; scale bar = 0.3 mm.

164 Dany Azar & André Nel / ZooKeys 130: 153-165 (2011)

Figure 10. Drawing of hypandrium of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male;
par = paraproct, ep = epiproct, trich = trichobothria; scale bar = 0.3 mm.

Figure | 1. Photograph of hypandrium and aedeagus of Libanopsyllipsocus alexanderasnitsyni gen. et sp.
n., holotype, male.

Oldest Psyllipsocidae from the Lebanese amber... 165

Figure | 2. Photograph of paraproct of Libanopsyllipsocus alexanderasnitsyni gen. et sp. n., holotype, male;

arrow shows the anal spine.